Cognitive Control - University of Amsterdam Psychology PDF

Cognitive control Gazzaniga | Ch. 12 Timo Stein www.timostein.net [email protected] What to study? Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A h...

Cognitive control Gazzaniga | Ch. 12 Timo Stein www.timostein.net [email protected] What to study? Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides)  Cognitive control – Overview Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides) Reference: p. 516–520  What to do next...? Input is often While output is limited to one, very complex... maybe two, actions at a time... Reference: p. 519–520  What to do next...? In every environment (input), we can act in But how does the brain choose among many different ways... these behavioral options (output)? Reference: p. 519–520  Habits What to do next...? Habits Action is automatically triggered by a stimulus or context Stimulus Response Outcome Not under voluntary control of a desired outcome/goal/reward Time Goal-directed action Goal-directed action Action to achieve a desired outcome/goal/reward Requires knowledge of a relationship between action and outcome Cognitive control is needed for Stimulus Response Outcome goals to influence behavior Time Reference: p. 519–520  Two prefrontal control systems 1 Goal-directed behavior Lateral prefrontal cortex (PFC), frontal pole Maintenance of goals in working memory Filtering of information according to goals, and goal-dependent initiation, inhibition, and shifting of behavior Planning and organization of multiple goals of different complexity 2 Conflict monitoring Medial PFC (incl. anterior cingulate cortex [ACC]) Monitoring of goal achievement (error detection, negative feedback, response conflict, surprise) Modulation of the degree and allocation of Reference:cognitive control p. 516–517, p. 564  Cognitive control deficits after PFC lesions No directly obvious impairments of perception, language, long-term memory, or motor skills. Complex, idiosyncratic impairments in goal-directed initiation, inhibition, and shifting of behavior (great difficulties in managing daily life), e.g.: Perseveration (persisting in a response even after being told that it is incorrect). Apathy, distractibility, impulsivity. Inability to make decisions, plan actions, understand consequences of actions, follow rules. Disregard social conventions, are socially inappropriate. Reference: p. 518  Environmental dependency syndrome Stimulus-driven behavior: actions guided not by the patient’s own goals but by what is available in the immediate surrounding environment Imitation behavior: for example, imitating the physician (hand gestures, body postures, drawing, combing hair, chewing on a pencil, speaking, singing, etc.) Utilization behavior: abnormal reliance on environmental stimuli to trigger behavior (e.g., repeatedly drinking from a glass without being thirsty) Reference: p. 518-519  Disproportionally large prefrontal cortex (PFC) in primates Reference: p. 517  Disproportionally large PFC in humans? Outdated idea by Brodmann (1912): Human PFC is disproportionally large compared to other non-human primates Reference: p. 517  Relative PFC size does not explain human cognition Only absolute PFC size is larger in humans than in non-human primates but proportions are similar across primates Reference: p. 517  But greater relative PFC white matter volume in humans Disproportionally greater expansion of white matter (axonal connections) in human PFC compared to other primates Reference: p. 517  Cognitive control – Overview Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides) Reference: p. 520–522  Two prefrontal control systems 1 Goal-directed behavior Lateral prefrontal cortex (PFC), frontal pole Maintenance of goals in working memory Filtering of information according to goals, and goal-dependent initiation, inhibition, and shifting of behavior Planning and organization of multiple goals of different complexity 2 Conflict monitoring Medial PFC (incl. anterior cingulate cortex [ACC]) Monitoring of goal achievement (error detection, negative feedback, response conflict, surprise) Modulation of the degree and allocation of Reference:cognitive control p. 516–517, p. 564  Delayed-response tasks Require to retain a stimulus attribute (e.g., location) not currently present in the environment ”in mind” over a delay period Performance on delayed-response tasks is impaired by PFC lesions In associative-memory control task, reward is paired with a cue (long-term memory association) Performance on the associative-memory control task is not impaired by PFC lesions Delayed-response tasks are widely used to study the neural bases of working memory Reference: p. 520–522  Delay-period activity Oculomotor delayed response (ODR) task: keep spatial location (cue) in mind to direct a later eye movement (response) Lateral PFC neurons fire continuously during delay (between cue and response) period for their preferred location Delay-period activity also for various other memorized stimulus attributes and other response modalities Delay period Reference: p. 520–522  Delay-period activity Successive memorization of two stimulus attributes: first identity, second location Some lateral PFC neurons retain identity information, others location information, and others both Task-specific selectivity of lateral PFC neurons Flexibility: when task changes, these neurons can retain information about different stimulus attributes Suggests that these neurons represent task goals rather than task-relevant information per se Reference: p. 520–522  PFC interactions with posterior cortex Integrative model of goal-directed processing Lateral PFC: sustained representation of task goal Posterior cortex (higher-level sensory areas): representation of task-relevant information (e.g., stimulus representations and knowledge in long-term memory) Reference: p. 522–524 Cognitive control – Key points I Why cognitive control? Prefrontal cortex For goals (rather than stimulus or context) Lesions: complex cognitive control deficits, to influence behavior e.g., perseveration, inability to plan goal- directed actions Know the difference between goal-directed action and habits (stimulus–response– Environmental dependency syndrome outcome) (imitation and utilization behavior) Disproportionally large PFC in primates Two cognitive control systems in PFC and/or humans? (gray vs. white matter) Lateral PFC/frontal pole: keeping goal in working memory, planning actions and Working memory attentional filtering according to goals Know details on delayed-response tasks and Medial PFC (incl. ACC): monitoring (error delay-period activity in lateral PFC and conflict detection, feedback and Lateral PFC sustains task goals in working surprise processing) and corresponding memory but interacts with posterior cortex modulation of the degree of cognitive containing task-relevant (stimulus) control representations (long-term memory)  Cognitive control – Overview Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides) Reference: p. 522–524, p. 540–542, p. 545–552  Two prefrontal control systems 1 Goal-directed behavior Lateral prefrontal cortex (PFC), frontal pole Maintenance of goals in working memory Filtering of information according to goals, and goal-dependent initiation, inhibition, and shifting of behavior Planning and organization of multiple goals of different complexity 2 Conflict monitoring Medial PFC (incl. anterior cingulate cortex [ACC]) Monitoring of goal achievement (error detection, negative feedback, response conflict, surprise) Modulation of the degree and allocation of Reference:cognitive control p. 516–517, p. 564  PFC interactions with posterior cortex Integrative model of goal-directed processing Lateral PFC: sustained representation of task goal Posterior cortex (higher-level sensory areas): representation of task-relevant information (e.g., stimulus representations and knowledge in long-term memory) Reference: p. 522–524  PFC interactions with posterior cortex Integrative model of goal-directed processing Lateral PFC: sustained representation of task goal Posterior cortex (higher-level sensory areas): representation of task-relevant information (e.g., stimulus representations and knowledge in long-term memory) Selection of task-relevant information: Dynamic filtering Lateral PFC: selection of different types of information according to dynamic goals (attentional mechanism) in posterior cortex Posterior cortex (higher-level sensory areas): inhibition of task-irrelevant and enhancement of task-relevant information Reference: p. 540–542  Dynamic filtering Attentional selection: Enhancement and suppression Task goals modulate posterior cortex: fMRI activity in category-selective higher-level visual cortex Category-specific enhancement of task-relevant information and suppression of task-irrelevant information (relative to passive viewing) Reference: p. 546–547  Dynamic filtering Attentional selection: Enhancement and suppression Task goals modulate posterior cortex: fMRI activity in category-selective higher-level visual cortex Category-specific enhancement of task-relevant information and suppression of task-irrelevant information (relative to passive viewing) Role of lateral PFC in attentional selection Filtering deficits with lateral PFC lesions: ERPs (P100) reveal reduced suppression of unattended tones and reduced enhancement of attended tones Reference: p. 545–546  Dynamic filtering Attentional selection: Enhancement and suppression Task goals modulate posterior cortex: fMRI activity in category-selective higher-level visual cortex Category-specific enhancement of task-relevant information and suppression of task-irrelevant information (relative to passive viewing) Role of lateral PFC in attentional selection Filtering deficits with lateral PFC lesions: ERPs (P100) reveal reduced suppression of unattended tones and reduced enhancement of attended tones Repetitive TMS (rTMS) to lateral PFC reduces attentional modulation of P100 in feature-based attention task Deficits primarily reflect reduced inhibition of Reference:irrelevant p. 547–548 information (e.g., larger P100 for  Inhibition of action Stop-signal task Respond to stimulus but inhibit response if a stop signal is presented shortly after the stimulus Stop signal typically activates (right) inferior frontal gyrus (similar for failed and successful stop) Reference: p. 549–550  Inhibition of action Stop-signal task Respond to stimulus but inhibit response if a stop signal is presented shortly after the stimulus Stop signal typically activates (right) inferior frontal gyrus (similar for failed and successful stop) Motor cortex: high initial activation level in failed stop trials (cannot be stopped by inferior frontal gyrus anymore) Similar to inferior frontal: preSMA in medial frontal cortex and basal ganglia (subthalamic nucleus STN) Aborting of response is carried out via pathway from inferior frontal to STN Reference: p. 549–550  Cognitive control – Overview Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides) Reference: p. 539–540, p. 542  Two prefrontal control systems 1 Goal-directed behavior Lateral prefrontal cortex (PFC), frontal pole Maintenance of goals in working memory Filtering of information according to goals, and goal-dependent initiation, inhibition, and shifting of behavior Planning and organization of multiple goals of different complexity 2 Conflict monitoring Medial PFC (incl. anterior cingulate cortex [ACC]) Monitoring of goal achievement (error detection, negative feedback, response conflict, surprise) Modulation of the degree and allocation of Reference:cognitive control p. 516–517, p. 564  Action hierarchy Making plans to organize actions A hierarchy of subgoals leading to a goal Planning: anticipation of consequences, requirements for achieving subgoals Impaired in patients with PFC lesions Reference: p. 539  Prefrontal cortex hierarchy Posterior-to anterior gradient in PFC related to the complexity/level of abstraction of action goals fMRI study with 4 nested tasks that increased in complexity/level of abstraction: A. Response competition: variation of the number of possible finger responses to colors (1–4) B. Feature task: response based on texture, colors indicate texture-response mapping C. Dimension task: colors indicate the dimension on which stimuli should be judged D. Context task: same as dimension task, but color- to-dimension mapping changes from block to block Reference: p. 539–540  Prefrontal cortex hierarchy Posterior-to anterior gradient in PFC related to the complexity/level of abstraction of action goals fMRI study with 4 nested tasks that increased in complexity/level of abstraction: A. Response competition: variation of the number of possible finger responses to colors (1–4) B. Feature task: response based on texture, colors indicate texture-response mapping C. Dimension task: colors indicate the dimension on which stimuli should be judged D. Context task: same as dimension task, but color- to-dimension mapping changes from block to block Shift from posterior premotor regions (A, B) to more anterior inferior frontal (C) and frontal pole Reference: p. 539–540  Cognitive control – Overview Introduction to cognitive control and prefrontal cortex (p. 516–520) Working memory in cognitive control (p. 520–522) Dynamic filtering and inhibition (p. 522–524, p. 540–542, p. 545–552) A hierarchy of goals in prefrontal cortex (p. 539–540, p. 542) Monitoring and the medial prefrontal cortex (p. 554–560, slides) Reference: p. 539–540, p. 542  Two prefrontal control systems 1 Goal-directed behavior Lateral prefrontal cortex (PFC), frontal pole Maintenance of goals in working memory Filtering of information according to goals, and goal-dependent initiation, inhibition, and shifting of behavior Planning and organization of multiple goals of different complexity 2 Conflict monitoring Medial PFC (incl. anterior cingulate cortex [ACC]) Monitoring of goal achievement (error detection, negative feedback, response conflict, surprise) Modulation of the degree and allocation of Reference:cognitive control p. 516–517, p. 564  Error detection and negative feedback ERP components linked to medial prefrontal cortex (ACC) Error-related negativity follows incorrect responses Feedback-related negativity follows feedback about errors Signals from ACC could be sent to lateral PFC to reactivate the goal in working memory Reference: p. 556–557  Error detection and negative feedback Predicting errors from fMRI activity Cognitive control network Activity in the cognitive control network (medial and lateral PFC): steady decrease before an error Reference: p. 557  Error detection and negative feedback Predicting errors from fMRI activity Cognitive control network Activity in the cognitive control network (medial and lateral PFC): steady decrease before an error Activity in the default mode network (precuneus, concerned with self-referential thinking): steady increase before an error Continuous shift from cognitive control to default mode network until an error is made (and then the shift reverses) Problems with the error detection Default mode hypothesis network Could partially reflect surprise (unexpected) ACC activity is also observed in tasks where errors are rare, but that induce conflict Reference: p. 557  Conflict monitoring Response conflict in the Stroop task Irrelevant words automatically evoke a response that can interfere with producing the correct ink name (in incongruent trials) Reference: p. 558–560  Conflict monitoring Response conflict in the Stroop task Irrelevant words automatically evoke a response that can interfere with producing the correct ink name (in incongruent trials) Incongruent trials (conflict) evoke activity in ACC ACC activity (and response times) are greater when the previous trial is congruent rather than incongruent (more conflict, more need for monitoring) Increased activity in ACC results in increased activity in lateral PFC in the next trial → Conflict monitoring reactivates the goal in working memory Reference: p. 558–560  Conflict monitoring ACC Posterior-to-anterior hierarchy in ACC Similar to posterior-to-anterior hierarchy in lateral prefrontal cortex In ACC control of conflict between: ▪ Posterior: potential motor responses ▪ Middle: possible response options Connectivit ▪ Anterior: possible response strategies y with ACC Topographical functional connectivity between ACC and lateral PFC in resting-state fMRI Suggests a specific signal from ACC to lateral PFC calling for greater goal activation Lateral PFC Reference: Slides only Cognitive control – Key points II Dynamic filtering and inhibition by lateral Monitoring in medial PFC (ACC) PFC Error detection and feedback processing: Lateral PFC can enhance and suppress Error-related and feedback-related representations in posterior cortex negativity ERPs from ACC according to task goals Shift from cognitive control to default mode Stop-signal task: inhibition of action network before an error involving lateral PFC plus motor areas Conflict monitoring in ACC with the Stroop- (know different response profiles of task example: Interplay between ACC and different areas) lateral PFC Action hierarchy → lateral PFC hierarchy Whole system dynamics: ACC detects conflict, signals to lateral PFC to reactivate Posterior-to-anterior gradient in lateral PFC reflecting complexity/level of the goal in working memory abstraction of action goals Similar hierarchy reflecting complexity of conflict monitoring in medial PFC (ACC)